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Goodyera repens

Table of Contents

• INTRODUCTORY
• DISTRIBUTION AND OCCURRENCE
• GENERAL INFORMATION ON BIOLOGY, ECOLOGY, FIRE, AND MANAGEMENT
• APPENDIX: FIRE REGIME TABLE
• REFERENCES

INTRODUCTORY

		FEIS ABBREVIATION: GOOREP COMMON NAMES: lesser rattlesnake plantain creeping rattlesnake plantain dwarf rattlesnake plantain northern rattlesnake plantain TAXONOMY: The scientific name of lesser rattlesnake plantain is Goodyera repens (L.) R. Br. (Orichidaceae) [28,31,47]. SYNONYMS: Goodyera repens var. ophiodes [30,66,84]. LIFE FORM: Forb
Photo used by permission of David McAdoo and the North Carolina Native Plant Society

DISTRIBUTION AND OCCURRENCE

North American distribution of lesser rattlesnake plantain. Map courtesy of the Flora of North America Association. 2013, 29 April.

Lesser rattlesnake plantain is native to North America [28] and Eurasia [87]. This review describes lesser rattlesnake plantain as it occurs in North America. Lesser rattlesnake plantain is transcontinental in Canada and occurs in continuous and discontinuous populations in the United States. Its northern range occurs in Alaska, Yukon, and Newfoundland. In the eastern United States, its southern range reaches North Carolina and Tennessee; in the West, it occurs in disjunct populations in Arizona, New Mexico, and Colorado [28].

States and provinces [78]:
United States: AK, AZ, CO, CT, ID, MA, MD, ME, MI, MN, MT, NC, NH, NM, NY, OH, PA, SD, TN, VA, VT, WI, WV, WY
Canada: AB, BC, LB, MB, NB, NF, NS, NT, ON, PE, QC, SK, YT

Site Characteristics:
Lesser rattlesnake plantain is associated with undisturbed, late-successional and old-growth forests. In much of its range, lesser rattlesnake plantain occurs in shady, moist, coniferous or mixed forests [2,11,17,28,31,40,53,57,69] on mossy or humus-covered ground [28,80], and can sometimes occur in bogs or swamps [28]. However, it also occurs in xeric conditions in the southern Appalachian Mountains including high, rocky, exposed cliffs and slopes of the Carolina hemlock (Tsuga caroliniana) community [77]; xeric heath balds and forest heathlands [56,86]; and relatively xeric eastern hemlock (Tsuga canadensis) forest [79,85].

Lesser rattlesnake plantain occurs at elevations ranging from sea level to about 9,500 feet (0-2,900 m) [28]. This varies with geographic location. For example, lesser rattlesnake plantain occurs at high elevations in disjunct populations in Colorado (8,065-8,258 feet (2,460-2,515 m)) [27] and New Mexico (7,450 feet (2,270 m)) [45] and occurs at low elevation in eastern Canada (450 feet (135 m)) [39].

Soils: Lesser rattlesnake plantain can tolerate a range in soil moisture regimes, from wet sites in the northern part of its range to dry sites in the southern Appalachians. In its northern range, lesser rattlesnake plantain typically grows on sites that have a thick organic layer or moss cover [2,43,63,69,76,80]. Soils on these sites tend to have a high water holding capacity or moderate drainage [34,53,69,83]. In Alaska, lesser rattlesnake plantain sometimes occurs on sites with permafrost [83]. In xeric coniferous heathlands of the southern Appalachian Mountains, lesser rattlesnake plantain occurs on dry, nutrient poor, leached, acidic sites and is classified as an acidophile [56,86]. Lesser rattlesnake plantain is considered an indicator species in serpentine woodlands of the southern Appalachian region [79].

Plant Communities:
Lesser rattlesnake plantain occurs in many vegetation types including late-successional coniferous forests [80,81,83], mixed hardwood forests [12,74], somewhat dry forests [31], heath balds and forest heaths [86], and bogs and swamps [28]. It is commonly associated with white spruce (Picea glauca) in boreal forests [22,49,53,76,80,81,83], and eastern hemlock in eastern montane forests [13,21,56,61,68,77,79,85].

Lesser rattlesnake plantain is an indicator of mature vegetation throughout its range. On the Tanana River, Alaska, lesser rattlesnake plantain is an indicator of the mature floodplain successional stage dominated by white spruce (150-200 years old) [32]. In southwestern Manitoba, lesser rattlesnake plantain was most common in mature northern whitecedar (Thuja occidentalis)-balsam fir (Abies balsamea)-black spruce (Picea mariana)-white spruce communities [37]. In boreal mixedwood stands in northern Ontario, lesser rattlesnake plantain was an indicator of unharvested forest [73]. In mature boreal mixedwood forest stands of western Canada, lesser rattlesnake plantain was an indicator of conifer forest patches (mostly dominated by white spruce) rather than broadleaf forest patches (mostly quaking aspen (Populus tremuloides)) (P=0.003) [82].

See Table 1 for additional descriptions of plant communities and site characteristics in areas where lesser rattlesnake plantain typically occurs or has been observed. See the Fire Regime Table for a list of plant communities in which lesser rattlesnake plantain may occur and information on the fire regimes associated with those communities.

GENERAL INFORMATION ON BIOLOGY, ECOLOGY, FIRE, AND MANAGEMENT

Lesser rattlesnake plantain leaves and inflorescence. Photos used by permission of David McAdoo and the North Carolina Native Plant Society.

Raunkiaer [67] life form:
Hemicryptophyte

Seasonal Development:
Lesser rattlesnake plantain is an evergreen perennial with leaves active throughout the winter [29]. The flowering period ranges from early July to mid-September depending on its location. In Montana, flowering begins in late July or early August and extends into mid-September [2]. In northern Michigan, flowering was observed from mid-July to late August with a median bloom date of July 31 [35]. In the mountains of Virginia, lesser rattlesnake plantain flowers in July and August [52]. After the plant blooms, the flowering rosette dies, and buds along the rhizome are released from dormancy [17].

Regeneration processes:
Lesser rattlesnake plantain regenerates sexually through seeds and vegetatively via rhizomes. Rhizomes that arise from seed require 5 years to produce a rosette of evergreen leaves, which produces a flowering stalk and seed capsules after approximately 3 more years [48]. The tiny seeds of lesser rattlesnake plantain are wind dispersed [2,5]. No information was available on lesser rattlesnake plantain soil seed banks in North America at the time of this writing (2013); however, research from old-growth Norway spruce (Picea abies) forests in Norway suggests that lesser rattlesnake plantain does not have a long-lived seed bank [71,72]. Goodyera species often form colonies due to their rhizomatous growth habit [17].

Interbreeding among Goodyera species is possible where mixed-species populations occur. See Kallunki [35] for details. Although lesser rattlesnake plantain is geitonogamous, self-pollination is unlikely to occur in nature because the flowers are slightly protandrous. When experimentally pollinated, lesser rattlesnake plantain collected from northern Michigan was capable of self-fertilization. Bumblebees are the principal pollinators of Goodyera species in North America [35].

Like other terrestrial orchids, lesser rattlesnake plantain requires a mycorrhizal symbiont for seed germination and early seedling development [4,10,24]. The symbiont is usually Rhizoctonia goodyerae-repentis. Lab experiments have demonstrated that lesser rattlesnake plantain can be germinated asymbiotically in the presence of an external carbohydrate source, whereas inoculated lesser rattlesnake plantain does not require a carbohydrate [24,65]. Other lab experiments with seeds collected from Pinnacle Mountain, South Carolina, found that germination rates were greater when seeds were exposed to light (37% germination) compared to those incubated in continuous darkness (25% germination), and light had no effect on seed mortality [55].

Successional status:
Lesser rattlesnake plantain is a shade-tolerant, mid- to late-successional species generally occurring in stands more than 70 years old [22,23,32,33,36,46,69,80,83].

In white spruce-fir and black spruce forests across the North American taiga from Newfoundland to Alaska, lesser rattlesnake plantain was prevalent in mature, undisturbed forest stands (i.e., stands that had no evidence of fire, insect infestation, or other major natural disturbance). Lesser rattlesnake plantain had a frequency of 59% in white spruce-fir stands and 54% in black spruce stands [39].

Chronosequence studies of primary succession on Alaskan floodplains indicate that lesser rattlesnake plantain typically occurs in mature white spruce stands that range from 120 to 300 years old [32,80,81,83]. On the Tanana River in interior Alaska, lesser rattlesnake plantain occurs in 150- to 300-year-old stands after balsam poplar-dominated stands shift to closed white spruce and develop a nearly complete ground cover of feathermosses (Hylocomiaceae) [32,80,81]. One of the studies on the Tanana River floodplain characterized lesser rattlesnake plantain as an indicator of the mature white spruce stage; it was not found in earlier successional stages or the subsequent black spruce stage [32]. Near the Chena River in interior Alaska, lesser rattlesnake plantain had 40% frequency in 120-year-old white spruce stands, but was absent from younger willow and balsam poplar stands and older (220-year-old) white spruce-black spruce and black spruce/Sphagnum stands [83]. Similarly, along the Peace River in British Columbia, lesser rattlesnake plantain is generally more common on sites dominated by white spruce, feather moss (Hylocomium splendens), and Schreber's moss (Pleurozium schreberi) and less common on sites dominated by earlier successional balsam poplar [76].

On boreal sites in eastern Canada, lesser rattlesnake plantain occurred only in mid- to late-successional stands [22,23]. Around Lake Duparquet in northwestern Quebec, lesser rattlesnake plantain appeared in stands dominated by balsam fir, black spruce, and paper birch (Betula papyrifera) 74 to 143 years after fire [22]. A study of mature second-growth stands and undisturbed forest stands at 2 stages of development (senescent and old growth) in the Gaspé Peninsula, Quebec, found that lesser rattlesnake plantain was not present in any early second-growth forests (50-year-old, even-aged balsam fir-white birch stands) but had 6% frequency in senescent forests (70-year-old, even-aged balsam fir stands) and 3.3% frequency in old-growth forests (90-year-old, uneven-aged balsam fir-black spruce and uneven-aged balsam fir-white spruce stands) [23]. In upland plantations and managed stands of jack pine (Pinus banksiana) and black spruce in Ontario, lesser rattlesnake plantain frequency corresponded with the greatest time since disturbance and stand establishment. Measurements taken over 20 years showed that lesser rattlesnake plantain increased from 6% to 11% mean relative frequency in jack pine stands, and from 4% to 17% mean relative frequency in black spruce stands [33].

On southern boreal mixedwood sites in Michigan, Manitoba, and Saskatchewan, lesser rattlesnake plantain is associated with late-successional, mixed coniferous forest [46,69]. In a study of primary succession across a sand-dune chronosequence bordering northern Lake Michigan, lesser rattlesnake plantain was found in trace amounts (<0.1% cover) on dune ridges that ranged from 175 to 835 years old. It was not present on the earliest successional dunes (25 years old) dominated by beach grass (Ammophila breviligulata), or on 55- to 175-year-old dunes dominated by evergreen shrubs. It was first present after the development of mixed pine forest (225-440 years old) consisting of eastern white pine (Pinus strobus), red pine (Pinus resinosa), white spruce, balsam fir, northern whitecedar, and paper birch [46]. In the mixedwoods of Manitoba and Saskatchewan, lesser rattlesnake plantain occurs after the development of the feathermoss stratum, which is rarely present until stands are more than 70 years old [69].

Lesser rattlesnake plantain is uncommon in Montana, but it sometimes occurs in older forests dominated by Douglas-fir with a well-developed moss layer dominated by feather moss and Schreber's moss [2], or on northerly aspects in late-successional spruce (Picea) or subalpine fir (Abies lasiocarpa) forests [63].

FIRE ECOLOGY:
Postfire regeneration strategy [75]:
Rhizomatous herb, rhizome in organic layer

Fire adaptations and plant response to fire:
Fire adaptations: Although no information was available on the immediate effects of fire on lesser rattlesnake plantain at the time of this writing (2013), most literature describes lesser rattlesnake plantain as poorly adapted to fire [2,63,70,88]. Its roots, like those of other Goodyera species, rarely penetrate the mineral soil [17]. Its rhizomes are shallowly rooted in mosses and the upper organic layer [2,63], which makes them susceptible to damage from fire. Habitat and germination/establishment requirements of lesser rattlesnake plantain (i.e., shade, thick organic layer, intact mycorrhizal community) make it unlikely to "respond well to fire" [2]. Rowe [70] characterizes lesser rattlesnake plantain as a fire avoider; these are defined as "shade-tolerant species that slowly reinvade burned areas; late successional, often with symbiotic requirements". Avoiders cannot establish immediately after fire but only after some habitat requirement has been met (e.g., sufficient shade or humus accumulation). These species occur late in succession in areas with long fire-return intervals [70].

Plant response to fire: Lesser rattlesnake plantain is consistently absent from postfire plant communities, suggesting that it is sensitive to fire. In a study on the effects of wildfire on understory plant community development in boreal mixedwood communities near Slave Lake, Alberta, lesser rattlesnake plantain was present in prefire sample plots but absent from postfire and manipulated (burned and sapling/herb layers removed) plots [49]. In northeastern Minnesota, lesser rattlesnake plantain had 20% and 13% frequency in unburned second-growth (about 70 years old), mixed conifer-hardwood stands but was nearly absent from 2 similar stands that were burned by wildfires. On 1 burned site, lesser rattlesnake plantain did not occur 3, 5, or 14 years after fire. On a second burned site, where "little or no soil burn occurred even though it was a hot fire", lesser rattlesnake plantain was not observed 3 or 5 years after fire and had a 3% frequency 11 years after fire [38]. Lesser rattlesnake plantain was absent from postfire communities in jack pine and black spruce plantations in northeastern Minnesota [3] and oak-pine (Quercus-Pinus) forest in North Carolina [25], despite its presence in similar unburned sites. In vegetation that was characterized as "standing virgin forest communities" before fire in northeastern Minnesota, only 1 individual lesser rattlesnake plantain was observed in the first 5 years after fire [62]; no information was given on prefire plant communities or unburned controls.

In the southern Appalachian Mountains in northeastern Tennessee, lesser rattlesnake plantain was observed in some postfire sites but at a lower frequency than in unburned controls. Zimmerman [88] examined the effects of prescribed fire on the herbaceous layer by comparing 6 burned sites with 6 unburned controls. Postfire measurements were taken between 2 and 6 years after fire. Leaf litter, canopy cover, soil moisture, and soil pH were "slightly affected" by the fire. Lesser rattlesnake plantain was absent from some of the 6 burned sites, whereas it was one of the most common herbaceous species in their unburned counterparts. Lesser rattlesnake plantain occurred 20 times in burned plots and 46 times in unburned controls. The research does not indicate whether the plants in burned sites occurred in unburned patches within the burn perimeter, were burned and sprouted after fire, or were newly germinated seedlings. The author suggests that lesser rattlesnake plantain "is an indicator of moist woods and is not adapted to a regular fire regime" [88].

On the Lewis and Clark National Forest in Montana, where lesser rattlesnake plantain is an imperiled species, stand-replacement fires (wildfire and prescribed) and timber harvests (clearcuts and seed tree cuts) may have eliminated or greatly reduced the occurrence of lesser rattlesnake plantain. One population may have been eliminated by the Turkey Fire, and 2 other populations were greatly reduced following stand-replacement wildfires. One of these wildfires left an area of unburned habitat where lesser rattlesnake plantain is still present. No prefire data on lesser rattlesnake plantain occurrence was available, but it likely occurred on these sites before the fire removed the shade and organic layers of the forest floor. Populations of lesser rattlesnake plantain extended up to the boundary of, but not into, a recent clearcut and severely burned forest in 2 river drainages of the Little Belt Mountains, suggesting that lesser rattlesnake plantain cannot withstand clearcutting and/or high severity fire [63].

Because lesser rattlesnake plantain occurs in a variety of communities, it is subject to many different fire regimes, but most of them have relatively long fire-return intervals. The Fire Regime Table summarizes characteristics of fire regimes for vegetation communities in which lesser rattlesnake plantain may occur. For additional information on fire regimes of communities where lesser rattlesnake plantain occurs, see FEIS reviews of dominant species such as white spruce and black spruce. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".

LEGAL STATUS AND MANAGEMENT:
Federal legal status:
None

Other status:
Information on state- and province-level protection status of plants in the United States and Canada is available at NatureServe. As of 1984, lesser rattlesnake plantain was ranked as apparently secure to vulnerable in Colorado, imperiled to vulnerable in Montana and North Carolina, imperiled in Arizona, Pennsylvania, and possibly Virginia, critically imperiled to imperiled in Kentucky and West Virginia, and critically imperiled in Massachusetts and Tennessee. It was possibly extirpated from Connecticut and Maryland [60]. Lesser rattlesnake plantain was ranked as apparently secure or secure in Canada, except in Nova Scotia where it is vulnerable and Prince Edward Island where it is imperiled [60].

Other management:
Lesser rattlesnake plantain is negatively affected by timber harvesting due to its need for shade and undisturbed forest floor. A Montana Field Guide [59] states that “any activities or events that substantially reduce the overstory and/or removes the duff/litter layer, such as thinning and logging operations or severe fires are likely to have adverse impacts” to lesser rattlesnake plantain. Lesser rattlesnake plantain is typically absent in the short to mid-term after all types of timber harvest. It was absent for at least 10 years after harvest in oak-hickory (Quercus-Carya) forests in southwestern Virginia and northeastern West Virginia [12]; it was observed 17 years after thinning and shelterwood harvest in the Lewis and Clark National Forest, Montana [63]; and it was absent from stands less than 70 years old after harvests in eastern boreal, balsam fir forest on the Gaspé Peninsula, Canada [23]. In uncut white spruce/alder communities near Fairbanks, Alaska, lesser rattlesnake plantain had 14% to 21% frequency but was absent during the first 2 years after logging. Of the 50 species present in the plots before logging, only 4 species–including lesser rattlesnake plantain–were absent after logging [26]. In the Lewis and Clark National Forest, Montana, lesser rattlesnake plantain was absent from a seed-tree harvest stand except in 2 small, undisturbed patches within the harvest unit. Abundant lesser rattlesnake plantain populations were observed in adjacent uncut stands. The author states that "it is likely that the loss of shade, organic matter, and moss during the clearcut, seed tree and dozer piling treatments, changed the microclimate on the forest floor to such an extent that lesser rattlesnake plantain could not survive or reproduce after treatment" [63].

APPENDIX: FIRE REGIME TABLE

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